Habitat linkages: the effect of intertidal saltmarshes and adjacent subtidal habitats on abundance, movement, and growth of an estuarine fish

In this study we used pinfish (Lagodon rhomboides) in field experiments to examine linkages between intertidal saltmarsh and adjacent subtidal habitats. Pinfish are more than twice as abundant in intertidal marshes adjacent to seagrass beds than in those adjacent to the unvegetated subtidal bottom. Movement of pinfish between the marsh edge and the adjacent subtidal habitat was greater for fish captured in areas with both intertidal and subtidal vegetation than in those with intertidal vegetation and adjacent unvegetated mudflats. This movement provides an important link between habitats, allowing transfer of marsh-derived secondary production to subtidal seagrass beds and vice versa. Pinfish held in enclosures with both intertidal and subtidal vegetation were, on average, approximately 90% heavier than fish held in enclosures with intertidal vegetation and unvegetated subtidal bottom. Because saltmarshes and seagrass beds contribute to the production of living marine resources, active measures are being taken to preserve and restore these habitats. The results from this study have direct application to decisions concerning site selection and optimal spatial proximity of saltmarsh and seagrass habitats in the planning of restoration and mitigation projects. To maximize secondary production and utilization of intertidal marshes, managers may opt to restore and/or preserve marshes adjacent to subtidal seagrass beds. Received: 31 May 1996 / Accepted: 23 October 1996     

Characterisation of microsatellite loci in the habitat-forming kelp, Ecklonia radiata (Phaeophyceae, Laminariales)

Habitat forming algae play an important role in the ecology of temperate reefs worldwide. Despite this, our understanding of levels of gene flow within and among populations of algae is largely limited to studies on intertidal species; we know comparatively little about important habitat-forming subtidal algae. Here, we develop eight polymorphic microsatellite markers for the characterisation of population genetic diversity and structure in the subtidal kelp, Ecklonia radiata. This large macroalga is the most abundant habitat-forming kelp on the subtidal rocky reefs of temperate Australia and New Zealand where it forms extensive forests that support an astounding diversity of associated taxa.     

First Record of Undaria Pinnatifida (Laminariales, Phaeophyta) in Southern Patagonia, Argentina

The Japanese kelp Undaria pinnatifida was found on 7th September 2005 in Ría Deseado (Santa Cruz, Argentina). This invasive algae was registered from the lower intertidal to the upper levels of the subtidal zone. The sporophytes were fixed to slabs, gravel and rocks of variable sizes and were also present as epibionts of tunicates. The sporophytes total length ranged between 3 and 88 cm, with a mean of 28.75 cm in the intertidal and 38.3 cm in the subtidal. The density and biomass increased from the intertidal to the subtidal. The kelp population showed a higher number of small immature individuals in the intertidal than␣subtidal where the organisms were bigger and with more individual biomass. This first record of U. pinnatifida in Southern Patagonia extends its southern distribution limit in Argentina, showing progressive expansion in the southwestern Atlantic. More surveys must be conducted to understand the evolution of U. pinnatifida invasion and its impact on the native benthic community.     

Facies architecture,depositional environments,and sequence stratigraphy of the Middle Cambrian Fasham and Deh-Sufiyan Formations in the central Alborz,Iran

Based on their lithologic characteristics and stratal geometries, the Middle Cambrian Fasham and Deh-Sufiyan Formations of the lower Mila Group in the Central Alborz, northern Iran, exhibit 39 lithofacies representing several supratidal to deep subtidal facies belts. The siliciclastic successions of the Fasham Formation are divided into two facies associations, suggesting deposition in a tide-dominated, open-mouthed estuarine setting. The mixed, predominantly carbonate successions of the Deh-Sufiyan Formation are grouped into ten facies associations. Four depositional zones are recognized on the Deh-Sufiyan ramp: basinal, outer ramp (deep subtidal associations), mid ramp (shallow subtidal to lower intertidal associations), and inner ramp (shoal and upper intertidal to supratidal associations). These facies associations are arranged in small-scale sedimentary cycles, i.e., peritidal, shallow subtidal, and deep subtidal cycles. These cycles reflect spatial differences in the reaction of the depositional system to small-scale relative sea-level changes. Small-scale cycles are stacked into medium-scale cycles that in turn are building blocks of large-scale cycles. Systematic changes in stacking pattern (cycle thickness, cycle type, and facies proportion) allow to reconstruct long-term changes in sea-level. Six large-scale cycles (S1–S6) have been identified and are interpreted as depositional sequences showing retrogradational (transgressive systems tract) and progradational (highstand systems tract) packages of facies associations. The six depositional sequences provide the basis for inter-regional sequence stratigraphic correlations and have been controlled by eustatic sea-level changes.     

Micropopulation differentiation in phenol content and susceptibility to herbivory in the Chilean kelp Lessonia nigrescenss (Phaeophyta,Laminariales)

Micropopulation differences in phenol content between intertidal and subtidal individuals of the kelp Lessonia nigrescens were found. Subtidal plants showed: (1) significantly higher phenol content than intertidal individuals, in vegetative and reproductive tissues, (2) intra-plant differences, with higher content in apical frond tissues, (3) higher resistance to consumption by herbivorous fishes. The microscopic progeny of subtidal plants showed the same trend as adult plants: (1) haploid spores from subtidal plants had higher phenol content than spores from intertidal individuals, and (2) the microscopic sporophytes derived from subtidal spores and gametophytes were less consumed by herbivorous snails (Tegula tridentata) than those derived from intertidal plant propagules. No increase in phenol content was detected after mechanical injury to experimental fronds, or after transplantation to the subtidal environment.In addition to the absence of inducible responses, the different phenol content between intertidal and subtidal individuals, in adult diploid plants and also in the haploid progeny, suggests that both environments differ someway enough to fix the mentioned features on the plants of Lessonia nigrescens. It is likely that the differences in herbivory between the two distributional extremes contributed to the observed pattern.     

A comparative analysis of the upper thermal tolerance limits of eastern Pacific porcelain crabs, genus Petrolisthes: influences of latitude, vertical zonation, acclimation, and phylogeny

Marine intertidal organisms are subjected to a variety of abiotic stresses, including aerial exposure and wide ranges of temperature. Intertidal species generally have higher thermal tolerance limits than do subtidal species, and tropical species have higher thermal tolerance limits than do temperate species. The adaptive significance of upper thermal tolerance limits of intertidal organisms, however, has not been examined within a comparative context. Here, we present a comparative analysis of the adaptive significance of upper thermal tolerance limits in 20 congeneric species of porcelain crabs, genus Petrolisthes, from intertidal and subtidal habitats throughout the eastern Pacific. Upper thermal tolerance limits are positively correlated with surface water temperatures and with maximal microhabitat temperatures. Analysis of phylogenetically independent contrasts (from a phylogenetic tree on the basis of the 16s rDNA gene sequence) suggests that upper thermal tolerance limits have evolved in response to maximal microhabitat temperatures. Upper thermal tolerance limits increased during thermal acclimation at elevated temperatures, the amount of increase being greater for subtidal than for intertidal species. This result suggests that the upper thermal tolerance limits of some intertidal species may be near current habitat temperature maxima, and global warming thus may affect the distribution limits of intertidal species to a greater extent than for subtidal species.     

Stratigraphic analysis of a fossil Neogoniolithon-capped patch reef and associated facies,San Salvador,Bahamas

An unusual Pleistocene patch reef is exposed in a coastal cliff at Grotto Beach, San Salvador, Bahamas. The reef is a coralline framestone constructed mainly by Porites astreoides together with a few large heads of Diploria strigosa and Montastrea annularis, and is capped by a dense thicket of Neogoniolithon strictum that is interpreted as marking the subtidal/intertidal boundary. The reef is flanked to the northeast by laminated to low-angle cross-laminated intraclastic grainstones and to the southwest by skeletal rudstone of reefal and interreefal derivation. Uranium-series dating of pure aragonite from a Diploria corallum yielded an age of 123 000±9000 years. Reef growth began on an erosional surface underlain by steeply crossbedded eolian grainstone. As the reef grew upward, it also grew laterally over adjacent penecontemporaneous subtidal sediments. The reef was eventually buried by 2.3 m of shallow subtidal and beach sediments that apparently prograded seaward during a highstand, or possibly while sea level was still rising. The shallow subtidal sediments are mainly peloidal, ooidal and skeletal grainstones that are pervasively bioturbated. The overlying beach facies comprises predominantly laminated, sparsely burrowed grainstone. The beach and shallow subtidal facies contain boulders of fine-grained laminated grainstone that are interpreted as storm-tossed blocks of beachrock. Living analogs of the Grotto Beach fossil reef lie off East Beach, San Salvador. Several of these have a flourishing cap of Neogoniolithon that extends above low-tide level and we believe that the Neogoniolithon cap of Grotto Beach reef did likewise. Wherever found in the stratigraphic record this facies should serve to identify the subtidal/intertidal boundary. The uppermost Pleistocene beach sediments associated with Grotto Beach fossil reef lie 5.8 m above present-day mean sea level, which ist strong evidence that this portion of San Salvador has undergone little subsidence since the Grotto Beach section was deposited.     

Intraspecific variation in the population dynamics and growth of the limpet,Cellana tramoserica

Summary Populations of the limpet Cellana tramoserica (Sowerby) from high, mid, and low intertidal regions, and from a subtidal zone, were studied at Cape Banks (N.S.W.), Australia. Individuals from the subtidal population had the largest mean and maximum shell size, the low and high shore populations were intermediate, while the midshore population had both the smallest mean and maximum size. The density of adults showed the reverse trend: the midshore region had the greatest adult population density while the subtidal population had the smallest density. The density of juveniles and recruits was negatively correlated with tidal height in the intertidal areas, but the density of both age-classes was smallest in the subtidal region. The rate of growth of individuals was negatively correlated to the adult density of a site; individuals from the subtidal population grew the fastest, while those from the midshore region grew the slowest. The level of adult mortality of the four populations was similar, with an annual rate of between 50–60%. Juvenile mortality did, however, differ among populations; reduced percentages of juveniles reached adult size in the mid and low shore populations than in the highshore and subtidal populations.The expected lifetime fecundity of individual females differed among the four populations. Subtidal individuals were expected to spawn 40 times the amount of gonad material during their life than were individuals from the midshore population. Individuals from the high and low populations probably spawn 10 times more than those from the midshore region. Differences in the fecundities of individuals were not likely to be the result of genetic differences because experimental transplants and manipulations of density showed that individuals from all the intertidal populations could increase their growth rate to match that of subtidal individuals. The implications of the difference in lifetime fecundity among populations are discussed.     

Production and population ecology ofPhyllospadix iwatensis Makino. II. Comparative studies on leaf characteristics,foliage structure and biomass change in an intertidal and subtidal zone

A seagrass in Japan,Phyllospadix iwatensis Makino, is distributed in the lower intertidal zone and upper subtidal zone making a dense population on the Choshi coast, Japan. IntertidalP. iwatensis is able to receive sufficient light for photosynthesis but experienced severe exposure to the air, which decreased a large amount of aboveground biomass in April to June (i.e. the daytime exposure season). SubtidalP. iwatensis was never exposed throughout the year and the aboveground biomass increased gradually over the daytime exposure season. However, the maximum aboveground biomass and shoot density of the subtidal plant never exceeded that of the intertidal plant. The dense foliage, large aboveground biomass and high shoot density of both intertidal and subtidal plants is likely to be an adaptation to heavy water movement, but the subtidal plants always received insufficient light for photosynthesis as a result of having dense foliage, particularly in turbid water. In choppy and swell sea,P. iwatensis did not seem to be adapted to growing in the subtidal zone where there was shortage of light.     

Pharmacology of scorpion (Lychas laevifrons Pock) venom with special reference to its neuromuscular activity

L. laevifrons venom caused irreversible blockade of electrically induced twitch responses on phrenic nerve diaphragm and chick biventer cervicis preparation. The venom lowered cat blood pressure, caused a brief cardiac arrest and increased cutaneous capillary permeability. It contracted several smooth muscle preparations. The quick contraction produced on guinea pig ileum was partly antagonized by mepyramine and completely by methysergide. The residual slow contraction was antagonized by SC 19220, a prostaglandin blocker. Haemolysis was not produced by the venom on human RBC. LD50 of crude venom in mice was 13.8 mg/kg (iv).     

A new late Katian (Late Ordovician) trilobite association from Zhenxiong,northeastern Yunnan,Southwest China and its palaeoecological implications

A new late Katian (Late Ordovician) trilobite association is documented from the Daduhe Formation in Zhenxiong, northeastern Yunnan, including 17 species of 15 genera, among which four species (Malongullia sinensis, M. zhenxiongensis, Taklamakania paucisegmentatus and Amphitryon constrictus) are new. Lithologic and faunal evidence enables the recognition of a new association with medium diversity, named the Taklamakania Association. It is a typical representative of the Raphiophorid Community that lived in a deep subtidal environment close to the anoxic basin. A total of five trilobite ecological associations were found to occur in South China during the late Katian, in relation to the environmental gradients ranging from shallow subtidal zone, deep subtidal zone, slope to dysoxic shaly basin.     

Site fidelity, home range, and tidal migrations of juvenile pinfish, Lagodon rhomboides, in salt marsh creeks

We conducted a series of experiments to determine the movements and fidelity of juvenile pinfish, Lagodon rhomboides, in salt marsh creeks within North Inlet Estuary, SC. In Experiment 1, we investigated the fidelity of pinfish (40–100mm SL) at four subtidal locations (9–617m apart) along the axis of a major creek. We trapped and marked 2,297 individuals and recaptured 15–22% of the fishes released at each site, almost all of which (99.7%) had been marked at the same location of recapture up to 3 months earlier. In Experiment 2, we investigated pinfish movements between subtidal and intertidal areas. In the 8 weeks prior to sampling 2 intertidal creeks, we marked 950 juveniles in the adjacent subtidal areas. Sampling of the 2 flooded intertidal creeks showed that 9–20% of the pinfish collected bore marks, and all had been previously marked at the subtidal site immediately adjacent to the intertidal creek. Gut analysis of 60 individuals revealed that juveniles collected from the intertidal areas at high tide had consumed about ten times more food than those collected at the subtidal sites at low tide. In Experiment 3, we determined the fidelity of recently settled pinfish (<35mm SL) and showed that 15% of the 434 marked pinfish remained in the same area; some were at liberty up to 7 weeks. In Experiment 4, we determined home range by tagging and recapturing juvenile pinfish at sites separated by 20m along a 200m subtidal transect. We estimated that during periods when the intertidal zone was not accessible, the average home range was 9.4m with only 10% of tagged fish moving greater than 20m during the 4 month study. Our results indicate that soon after recruitment to the estuary, pinfish establish strong fidelity for sites within salt marsh creeks and exhibit tidal periodicity in both movements and feeding.     

Movement,morphology, reproductive periodicity,and some factors affecting gonad production in the seastar Astrostole scabra (Hutton)

At Kaikoura, New Zealand (42°25′S: 173°42′E), tagged, immature Astrostole scabra (Hutton) usually remained within intertidal pools for <30 days. There was no seasonal vertical migration or intermixing of sub-populations. Intertidal seastars were smaller, had a thicker skeleton, and were damaged more frequently than their subtidal counterparts.The onset of sexual maturity in this species occurred coincidentally with the adoption of an exclusively subtidal habitat (20–146 m), the attainment of a radius of at least 110 mm, and a reduction in skeleton thickness. In 1975, 1976, and 1977, spawning was in late August-early September. Gametogenesis commenced in February–March. There was no seasonal inverse correlation between gonad and pyloric caeca indices.In undamaged, subtidal seastars of both sexes, gonad production was related to body size such that the volume of gonad produced by August increased linearly with respect to eviscerated wet weight. Regeneration partly suppressed gonad production, and large intertidal seastars had reduced gonad indices. The periodicity of reproductive events could not be correlated unequivocally with any physical or biotic exogenous environmental factor.     

A comparative study of territoriality in intertidal and subtidal blennioids (Teleostei, Blennioidei)

This paper presents a comparative analysis of territoriality in three intertidal (Lipophrys pholis, Coryphoblennius galerita, Salaria pavo) and two subtidal (Tripterygion delaisi, Parablennius pilicornis) blennioid fishes. Focal-animal observations of males guarding eggs showed that: (i) intertidal species had smaller territories that were less frequently patrolled; (ii) in subtidal species feeding was limited to the territory, while in intertidal fishes a substantial proportion of the feeding acts occurred outside the defended area; (iii) intertidal species spent less time out of the nest and showed a lower level of locomotory activity; (iv) subtidal species were subjected to a higher number of territorial intrusions with more species intruding their territory, some of them potential egg predators; (v) subtidal species performed several water column displays that played a significant role in courtship, while in intertidal fishes these displays are almost absent. Signalling without the presence of a female was almost absent in intertidal species. It is argued that intertidal fishes minimize the time spent out of the nest and the loss of contact with the substrate, features that may be of high survival value in conditions of strong turbulence. Although intertidal species have a reduced time available for feeding, this may be compensated by lower levels of locomotory activity, territorial defense and risk of egg predation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Relationship between algae and environment: an Early Cretaceous case study, Trascău Mountains, Romania

The relationship between algae and depositional environment was studied in a limestone succession of Berriasian–Valanginian age. Several depositional environments were recorded from shallow subtidal to intertidal and supratidal, with salinity ranging from normal marine to fresh and/or supersaline water. The algal assemblages consist mainly of dasycladaleans, rivulariacean-type cyanobacteria and charophytes. Nipponophycus (Bryopsydales) and Lithocodium-Bacinella (microproblematicum), occur at some levels. Dasycladaleans are associated with subtidal, sometimes restricted (“lagoonal”) environments, while rivulariacean-like cyanobacteria and charophytes characterise intertidal-supratidal and fresh and/or supersaline environments, respectively. Among the dasycladaleans, Salpingoporella annulata is often related to restricted environments where it forms monospecific assemblages. Large dasycladaleans, such as Selliporella neocomiensis, Macroporella praturloni and Pseudocymopolia jurassica are found in subtidal high-energy deposits (bioclastic grainstones). The relationship between environment and algae, characteristic for each depositional unit, can be used to interpret the relative sea-level variations.     

Sequence stratigraphy and carbonate platform organization of the Devonian Santa Lucia Formation, Cantabrian Mountains, NW-Spain

In this paper, the sedimentology and the stratigraphic architecture of the Devonian Santa Lucia Formation in the Cantabrian Mountains of NW-Spain are described. The Santa Lucia Formation consists of 11 different facies that can be attributed to peritidal/lagoonal, intertidal and subtidal facies associations. These facies associations are arranged in small-scale sedimentary cycles. Three different settings of small-scale sedimentary cycles are recognized: intertidal/supratidal, shallow subtidal/intertidal and subtidal cycles. These cycles reflect spatial differences in the reaction of the depositional system to small-scale relative sea-level changes. Small-scale stratigraphic cycles are stacked into seven medium-scale cycles that in turn are integral parts of three larger-scale cycles. Most of the Santa Lucia Formation (sequences 2–6) forms one major large-scale cycle, whereas sequences 1 and 7 are part of an underlying and an overlying cycle, respectively. Eustatic sea-level changes exerted major control on the formation of these large-scale sequences, whereas the medium-scale cycles seem to be co-controlled by regional tectonism and eustasy. Small-scale cycles seem to be the product of high frequency, eustatic sea-level changes. During the deposition of the Santa Lucia Formation, the morphology of the carbonate platform changed from a gently south-dipping ramp to a rimmed shelf and back to a gently dipping ramp.     

Long-term studies of macrozoobenthos in intertidal and shallow subtidal habitats near the island of Norderney (East Frisian coast,Germany)

Near the East Frisian island of Norderney two sites are investigated permanently in order to study long-term fluctuations of macrozoobenthos: one transect (since 1977) at the northern side in shallow subtidal waters and another one (since 1976) at the sheltered southern side in the intertidal area of the Wadden Sea. Since 1980 the investigations have been continued in the frame of COST-47, sedimentary intertidal programme (including the shallow subtidal habitats colonized by the Macoma balthica community).The results up to 1984, respectively to 1985, are presented and the changes of abundance of the dominant species are discussed. Discussed are also the influences of water temperatures and sediment disturbances caused by wave action. There is evidence that the intertidal variety of the Macoma balthica community shows a greater stability than the subtidal variety.     

Microbial-Activated Sediment Traps Associated with Oncolite Formation Along a Peritidal Beach,Northern Arabian (Persian) Gulf,Kuwait

Identification of microbial communities within shoreline sediments and sediment precipitates from the Tigris-Euphrates delta (northern Kuwait) were determined by microscopic/nanoscopic studies, and by molecular analysis. Oncolites are syn-diagenetic carbonate precipitates that are surviving in a shallow subtidal to intertidal siliciclastic environment with periodically excessive hydraulic energy, extreme salinity (up to 47 per mil), and high concentrations of organic matter. X-ray diffraction techniques reveal that oncolite cortices are predominantly composed of calcite, quartz, halite and dolomite, associated with minor fractions of clay minerals. Quantitative analysis of the Corey Shape Factor reveals distinct morphological populations but with local overlap. A plot of the Equivalent Diameter vs. Corey Shape Factor provided the best indicator of the morphological relationships within the total oncolite population, indicating a hydrodynamically controlled morphological distribution defining intertidal and subtidal oncolite classes. Direct microscopic examination of the samples indicates that diatoms are the most abundant eukaryotic algae in subtidal sediments and within actively precipitating carbonate cements, especially the genus Navicula. In contrast, filamentous cyanobacteria from the genus Anabaena are most abundant in the intertidal zone sediments. The PCR-DGGE of the 16SrRNA gene of the cyanobacteria shows a higher diversity for this genus of bacteria in all sediment samples and that the cyanobacterial population in the diagenetically precipitating oncolites are closely related to the population found in the subtidal sediments. Dunaliella viridis dominates the culturable algae obtained from the four tidal zones. Our results indicate that a range of microbial populations are actively contributing to the formation of microbially-induced sedimentary structures in the extreme conditions of the southern Tigris-Euphrates delta.     

The composition,structure, and distribution of benthic ostracod fauna (Ostracoda,Myodocopa) in Antarctic waters

The benthic fauna of ostracods of the order Myodocopida of Antarctic waters is characterized by high diversity, relative species abundance, and a complicated taxonomic and ecological structure, with a simplified biogeographical structure. This fauna, which is distinguished by a high level of endemicity, although at a low taxonomic rank, includes a great share of deep-sea and subtidal elements. Ostracod populations of High and Low-Antarctic subzones differ qualitatively and quantitatively. A distinct impoverishment of fauna is observed in the region of the Antarctic divergence compared to the more northern areas. The number of species increases with depth to reach its maximum in the lower subtidal zone and on the upper continental slope at depths of 200–500 m. The number of species decreases with increasing depth. Myodocopida have not been yet found in the Antarctic waters deeper than 5000 m.     

Canopy interactions and physical stress gradients in subtidal communities

Species interactions are integral drivers of community structure and can change from competitive to facilitative with increasing environmental stress. In subtidal marine ecosystems, however, interactions along physical stress gradients have seldom been tested. We observed seaweed canopy interactions across depth and latitudinal gradients to test whether light and temperature stress structured interaction patterns. We also quantified interspecific and intraspecific interactions among nine subtidal canopy seaweed species across three continents to examine the general nature of interactions in subtidal systems under low consumer pressure. We reveal that positive and neutral interactions are widespread throughout global seaweed communities and the nature of interactions can change from competitive to facilitative with increasing light stress in shallow marine systems. These findings provide support for the stress gradient hypothesis within subtidal seaweed communities and highlight the importance of canopy interactions for the maintenance of subtidal marine habitats experiencing environmental stress.